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Home , Asterids, Begoniaceae, Brachyglottis, Brachyglottis repanda, Eucalyptus leucophylla, Eucalyptus seeana

International Journal of Science, Environment ISSN 2278-3687 (O) and Technology, Vol. 7, No 5, 2018, 1786 – 1793 2277-663X (P)

DIVERSITY IN HOST PREFERENCE OF ROTYLENCHUS SPP. Y.S. Rathore Principal Scientist (Retd.), Indian Institute of Pulses Research, Kanpur -208024 (U.P.) E-mail: [email protected]

Abstract: of the Rotylenchus are ecto- or semi-endo parasites and feed on roots of their host . In the study it was found that 50% species of Rotylenchus were monophagous and mostly on plants in the followed by monocots, and gymnosperms. In general, Rosids and Asterids combined parasitized more than 50% host species followed by monocots. Though food preference was species specific but by and large woody plants were preferred from very primitive families like Magnoliaceae and Lauraceae to representatives of advanced families. Woody plants like pines and others made a substantial contribution in the host range of Rotylenchus. Maximum number of Rotylenchus species harboured plants in families Poaceae (monocots), (Rosids) and (Asterids) followed by , , and Pinaceae. It is, therefore, suggested that agricultural crops should be grown far away from wild vegetation and forest plantations. Keywords: Rotylenchus, , Rosids, Asterids, Gymnosperms, Host preference.

INTRODUCTION Species of the genus Rotylenchus (Nematoda: Haplolaimidae) are migratory ectoparasites and browse on the surface of roots. The damage caused by them is usually limited to necrosis of penetrated cells (1). However, species with longer stylet penetrate to tissues more deeply and killing more cells and called as semi-endoparasites (2,3). The genus contains 97 nominal species which parasitize on a wide range of wild and cultivated plants worldwide (3). Studies have revealed that generally nematodes prefer to parasitize host plants from selective taxonomic groups or some similar nature (4,5,6,7). It was, therefore, thought to be appropriate to find out the host preference of Rotylenchus species in relation to taxonomic groups of species. MATERIALS AND METHODS Host plants of Rotylenchus species were downloaded from Nemaplex (8) and other reported in the literature were added. All the plant species were aligned and classified according to APG IV system(9)(Fourth version of modern, mostly molecular based system of plant of flowering plants developed by Angiosperm Phylogeny Group). Here families are grouped into different . Host species were numerically categorized in each clade. Received Sep 23, 2018 * Published Oct 2, 2018 * www.ijset.net 1787 Y.S. Rathore

To observe the superiority of one clade over the other or one species over the other, Specific Affiliation Index (SAI) and General Affiliation Index (GAI) were computed following the method of Rathore and Tiwari (5). Certain ratios viz., species: genus, species: and species: were calculated to observe the contribution of particular group(s) towards the preference of host species. The terms monophagous, oligophagous and polyphagous were adopted from Barneys and Chapman (10). RESULTS AND DISCUSSION A general bird-eye picture evolved from Table 1 revealed that majority of plants were preferred from Rosids, Asterids and monocot clades from angiosperms followed by gymnosperms. Rosids and Asterid clades had 29.371 and 28.671% plants, respectively followed by monocots (18.881%). Rosids and Asterids combination resulted in more than 50% plants of the total. Although, Rotylenchus species parasitized very primitive Magnoliid plants (2.098%) and gymnosperms (11.189%). The Specific Association Index (SAI) did not indicate better preference between prominent clades. Their values demonstrate high polyphagy and wide distribution of host species in genera, families and orders. Contribution of aforesaid parameters was better in gymnosperms as evident by species: family and species: order ratios which were 2.000 and 2.667, respectively. In general, gymnosperms were preferred better than angiosperms in spite of angiosperms had bigger host range. Table 1. Association of Rotylenchus spp. to different host parameters Taxonomic Host Genera Families Orders SAI clades species Magnoliids 3 ( 2.098)* 2 (1.550) 2 (1.942) 2 (2.174) 0.833 26 19 17 Monocots 27 (18.881) 0.468 (20.155) (18.447) (18.498) 3 (2.098) 3 (2.326) 3 (2.913) 3 (3.261) 0.556 Superrosids 3 (2.098) 3 (2.326) 3 (2.913) 3 (3.261) 0.556 42 36 30 28 Rosids 0.468 (29.371) (27.907) (29.126) (30.435) 6 (4.196) 5 (3.876) 5 ( 4.854) 3 (3.261) 0.615 41 40 31 28 Asterids 0.434 (28.671) (31.008) (30.097) (30.435) COM Clade 2 (1.399) 2 (1.550) 2 (1.942) 2 (2.174) 0.667 16 Gymnosperms 12 (9.302) 8 (7.767) 6 (6.522) 0.692 (11.189) *Figures in parentheses are percent values

Host plants of individual Rotylenchus species are listed in Table 2. They are grouped in each clade along with the families. It was observed that out of 28 Rotylenchus species, 50% were Diversity in Host Preference of ROTYLENCHUS SPP. 1788

Table 2. Host taxonomic affinity of Rotylenchus spp. S.No. Rotylenchus Host species with taxonomic groups No. GAI Status spp . host species 1 R. agnetis Mono -Aspagaraceae (1) Ruscus aculentus 1 1 Monophagous 2 R. boocki Mono -Araceae (1) Anthurium andraeanum 1 1 Monophagous 3 R. breviglans Magnoliids -Lauraceae (1) Persea americana ; 4 0.6 Polyphagous Superrosids -Grossulariaceae (1) speciosum ; Superasterids -Cactaceae (1) Opuntia spp.; Asterids - (1) leucophylla 4 R. buxophilus Mono -Asparagaceae (3) Cordyline sp., Ruscus 28 0.448 Polyphagous hypophyllum, Sansevieri a sp.; Araceae (1) Anthurium sp., Arecaceae (1) Palmaceae sp.; Poaceae (1) Secale cereale; Strelitziaceae (1) Streltzia sp. Eudicots -Berberidaceae (1) Berberis thunbergii ; Buxaceae (1) Buxus sempervirens; Superrossids -Paeoniaceae (1) Paeonia sp.; Rosids- (1) sp.; Fabaceae (1) lunatus; (1) ; (3) sp., Eucalyptus terecornis, guajava ; Rosaceae (3) x ananasa, , sp .; Superasterids - Cactaceae (1) Cactaceae sp.; Asterids - (1) majo r; (1) arborea; Oleaceae (2) Ligustrum sp., ; (1) lycopersicum ; (1) Camelli a sp.; COM clade- (1) sp.; Gymnosperms -Pteridaceae (1) Adiantum sp. 5 R. Rosids -Fagaceae (2) , Quercus 2 1.333 Monophagous cazorlaensis rotundifolia 6 R. cretensis Asterids -Oleaceae (1) sp. 1 1 Monophagous 7 R. cypriensis Asterids -Oleaceae (1) Olea sp.; Solanaceae (1) 2 0.667 Polyphagous Solanum tuberosum 8 R. Mono -Asparagaceae (1) Ruscus hyrcanus 1 1 Monophagous dalikhaniensis 9 R. eximius Rosids -Rosaceae (1) dulcis ; (1) 3 0.556 Polyphagous Pistacea lentiscu s; Asterids -Oleaceae (1) Olea europaea sp . sylvistris 10 R. goodeyi Mono -Iridaceae (1) Iris sp.; Rosids -Fabaceae (1) 8 0.588 Polyphagous sativa ; Moraceae (1) Ficus sp.; Rosaceae (2) , Fragaria x ananasa ; Asterids -Asteraceae (1) ; Oleaceae (1) Olea europaea sp. sylvestris ; COM clad e- (1) sp. 11 R. graecus Mono -Poaceae (1) Arundo donax ; Asterids - 2 0.883 Polyphagous (1) helix 12 R. helenae Mono -Poaceae (1) Triticum aestivum ; Asterids - 2 0.883 Polyphagous Ericaceae (1) 13 R. incognitus Gymnosperms -Pinaceae (1) Pinus pinaster 1 1 Monophagous 14 R. incullus Rosids - (1) vinifera 1 1 Monophagous 15 R. iranicus Rosids -Fagaceae (1) 1 1 Monophagous 16 R. jaeni Rosids -Fagaceae (1) 1 1 Monophagous 1789 Y.S. Rathore

17 R. Mono -Poaceae (2) Festuca pseudovina, Zea may s; 3 0.719 Polyphagous jagatpurensis Asterids -Asteraceae (1) Artemissia maritima 18 R. labiodiscus Asterids -Asteraceae (2) repanda, 2 0.667 oligophagous Cyathia sp. 19 R. landii Gymnosperms -Pinaceae (1) Pinus pinaster 1 1 Monophagous 20 R. laurentinus Asterids - (1) carota 1 1 Monophagous 21 R. magnus Rosids -Fagaceae (1) Quercus suber ; Asterid s- 2 0.667 Polyphagous Aquifoliaceae (1) 22 R. montanus Rosids -Rosaceae (1) domesticus 1 1 Monophagous 23 R. pumilus Mono -Poaceae (1) Poa pratensis ; Gymnosperm s- 4 0.75 Polyphagous Cupressaceae (1) Juniperus monosperma ; Pinaceae (2) Pinus edulis, Pinus pinaster 24 R. robustus Mono -Alstroemeriaceae (1) Alstroemeria sp.; 59 0.545 Polyphagous Araceae (2) Zantedes chia aethiopica, Philodendron sp.; Arecaceae (2) Washingtonia sp., Cocos nucifera ; Asparagaceae (2) Convallaria majalis, Lilium speciosum ; Bromeliaceae (1) Ananas comosu s; Iridaceae (1) Gladiolus sp.; Poaceae (2) Lolium perenne, Saccharum officinarum; Eudicots- Buxaceae (1) Buxus semperviren s; Superrossids - (1) Huachera sp.; Rosids -Fabaceae (1) sativum; (1) sp.; (1) sp.; Myrtaceae (2) Eucalyptus sp., ; Rosaceae (3 ) Fragaria sp., , Rosa sp.; (2) sinensis , Citrus sp.; Vitaceae (1) ; Superasterids -Amaranthaceae (1) Beta vulgaris ; Cactaceae (1) Cactaceae sp.; Caryophyllaceae (2) Dianthus barbatus, Dianthus caryophyllus ;Asterids - (1) carlesi i; Apiaceae (1) ; Aquifoliaceae (1) ; Asteraceae (4) sp., sp., hispanica, Senec io sp.; (1) florida ; Ericaceae (4) Andr - omeda sp. , sp., japonica , sp.; (1) sp.; Oleaceae (2) Ligustrum sp., Olea europaea ; (1) sp.; Solanaceae (3) Solanum lycopersicum, Solanum tuberosum, tabacum; Theaceae (1) sp.; Magnoliids -Magnoliaceae (1) Magnoliax soulangian a; Gymnosperm s-Cupressaceae (4) Thuja occidentalis, Juniper sp., Juniper squamata, Sequoia sp.; Pinaceae (5) Picea pungens, Picea sitchensis, Pinus sp ., Pinus sylvestris, Pinus radiata; Pteridaceae (1) Adiantum pedatum; 25 R. uniformis Magnoliids -Lauraceae (1) Persea americana; 7 0.474 Polyphagous Rosids-Fabaceae (1) max ; Rosaceae (2) Malus sylvestris, Prunus domesticus; Sapindaceae (1) sp.; Asterids-Apiaceae (1 ) Daucus carota; Theaceae (1) 26 R. unisexus Rosids -Fabaceae (1) Glycine max ; Rutaceae (1) 2 0.667 Polyphagous Citrus aurantium 27 R. urmiaensis Rosids - (1) Corylus sp. 1 1 Monophagous 28 R. vitis Rosids -Vitaceae (1) Vitis vinifera 1 1 Monophagous Diversity in Host Preference of ROTYLENCHUS SPP. 1790

monophagous ( R. agnetis, R. boocki, R. cajorlaensis, R. cretensis, R. dalikhaniensis, R. incognitus,R. incullus, R. iranicus, R. jaeni, R. landii, R. laurentinus, R. montanus, R. urmiaensis, R. vitis ) and one species R. labiodiscus showed oligophagy (3.871%). Rest of the Rotylenchus species (46.429%) were polyphagous in nature. Large number of monophagous species hosted Rosids (25%) followed by monocots (10.714%), Asterids (7.143%) and gymnosperms (7.143%). This shows that host plants in Rosids have been comparatively of recent origin. Among polyphagous species, R. buxophilus and R. robustus had wide host range having 28 and 59 species, respectively representing many clades. General Affiliation Index (GAI) was 1.000 for all the monophagous species and the same was less than 1 for polyphagous. Hutchinson (11) divided dicotylednous angiosperms into two divisions viz. lignosae (trees and and some herbs clearly derived from and related to other woody plants – a fundamentally woody group) and herbaceae (herbs and rarely shrubby plants related to and derived from herbaceous stocks – a fundamentally herbaceous group). Since more than 50% host plants fall in Rosids and Asterids clades, and if we compare the families of these host species by placing them in lignosae (Apocynaceae, Araliaceae, Aquifoliaceae, Begoniaceae, Betulaceae, Cornaceae, Ericaceae, Fabaceae, Fagaceae, Hydrangeaceae, Malvaceae, Moraceae, Myrtaceae, Oleaceae, Rosaceae, Rutaceae, Sapindaceae, Theaceae, Vitaceae) and herbaceae (Apiaceae, Asteraceae, Adoxaceae, Geraniaceae, Lamiaceae, Plantaginaceae, Solanaceae) we observed that out of 26 families of Rosids and Asterids – 19 grouped in lignosae and 7 herbaceae. This is enough to indicate that Rotylenchus species have greater preference to woody plants. This view gets further support from the plant species belong to Magnoliids, which is entirely a woody group and perhaps the most ancient type of existing. In addition, families Grossulariaceae in Superrosids and Cactaceae in Superasterids were also classified in lignosae by Hutchinson (11) having woody plants. Gymnosperms include timber trees and all are woody in nature. Although the preference is species specific but in general there is strong affinity for host plants of woody nature. It is therefore necessary that agricultural crops should be grown away from forest areas and intercropping of only non-host species is suggested. Perusal of Table 3 revealed that family Poaceae in monocots, Rosaceae in Rosids and Oleaceae in Asterids harboured maximum number of nematode species having 6 in each case followed by Fabaceae (5), Fagaceae (4), Asteraceae (4) and Pinaceae (4), etc. R. robustus was the most prolific feeder and parasitized host species from 33 families and from very primitive to advanced ones. 1791 Y.S. Rathore

Table 3. Distribution of Rotylenchus spp. to host families and orders Clade Order Family Rotylenchus species No. of spp. Magnoliids Lauraceae R. breviglans, R. uniformis 2 Magnoliaceae R. robustus 1 Monocots Araceae R. boocki, R. buxophilus, R. robustus 3 Arecaceae R.buxophilus, R. robustus 2 Asparagaceae R. agnetis, R. buxophilus, R. 4 dalikhaniensis, R. robustus Iridaceae R. goodeyi, R. robustus 2 Bromeliaceae R. robustus 1 Poaceae R. buxophilus, R. graecus, R. helenae, R. 6 jagatpurensis, R.pumilus, R. robustus Strelitziaceae R. buxophilus 1 Eudicots Buxaceae R. buxophilus, R. robustus 2 Ranunculaceae R. buxophilus 1 Superrosids Grossulariaceae R. breviglans 1 Paeoniaceae R. buxophilus 1 Saxifragaceae R. robusts 1 Rosids Fabaceae R. buxophilus, R. goodeyi, R. robustus, 5 R. uniformis,R. unisexus Betulaceae R. urmiaensis 1 Fagaceae R. cazorlaensis, R. iranicus, R. jaeni, R. 4 magnus Geraniaceae R. robusts 1 Malvaceae R. robusts 1 Myrtaceae R. buxophilus, R. robustus 2 Moraceae R. buxophilus, R. goodeyi 2 Rosaceae R. buxophilus, R. eximius, R. goodeyi, R. 6 montanus,R. robusts, R. uniformis Rutaceae R. robustus, R. unisexus 2 Sapindaceae R. uniformis 1 Vitlaes Vitaceae R. incullus, R. robustus, R. vitis 3 Superasterids Amaranthaceae R. robustus 1 Cactaceae R. breviglans, R. buxophilus, R. robustus 3 Caryophyllaceae R. robustus 1 Asterids Apiaceae R. laurentinus, R. robustus, R. uniformis 3 Araliceae R. graecus 1 Asteraceae R. goodeyi, R. jagatpurensis, R. 4 labiodiscus, R. robustus Aquifoliaceae R. magnus, R. robustus 2 Cornaceae R. robustus 1 Hydrangeaceae R, robustus 1 Adoxaceae R. robustus 1 Ericaceae R. buxophilus, R. helenae, R. robustus 3 Theaceae R. buxophilus, R. robustus, R. uniformis 3 Apocynaceae R. buxophilus 1 Lamiaceae R. breviglans 1 Diversity in Host Preference of ROTYLENCHUS SPP. 1792

Oleaceae R. buxophilus, R. cretensis, R. cypriensis, 6 R. eximius,R. goodeyi, R. robusts Plantaginaceae R. robustus 1 Solanaceae R. buxophilus, R. cypriensis, R. robustus 3 COM clade Celastraceae R. buxophilus 1 Violaceae R. goodeyi 1 Gymnosperms Pinales Cupressaceae R. pumilus, R. robustus 2 Pinaceae R. incognitus, R. landii, R. pumilus, R. 4 robusus Polypodiales Pteridaceae R. buxophilus, R. robustus 2

Molecular studies of Subbotin et al (12) suggested that R. Goodeyi and R. laurentinus are very closely related or co-specific taxa. Volvas et al (1) reported that nematode presently known as R. Magnus magnus and R. Magnus jaeni are separate and distinguishable. Tzortzakakis et al (13) observed close molecular relationships between R. cretensis, R. urmiensis and R. striaticeps but all are separate species. In our study all are differing in their host preference. Acknowledgements Authors are grateful to Mr. Akhilesh Rathore for extending help in retrieving the information. References [1] Volvas, N., Subboti, S.A., Troccoli, A., Liebanas, G. and Castillo, P. (2008) Description of Rotylenchus montanus s. n. and recognition of R. jaeni comb. N. as a separate species with approaches to molecular phylogeny of the genus Rotylenchus (Nematoda, Tylenchida). Zoologicasoupta 37: 521-537. (doi: 10.1111/j 1463-6409.00337.x) [2] Decraemer, W. and Hunt, D.J. (2006) Structure and classification in Perry, R.N. and Moens, M. (Eds.) Plant nematology. CABI publishing, Oxfordshire, U.K., pp 3-32. (http://dx.doi.org/10.1079/9781845930561.0003) [3] Talezari, A., Pourjam, E., Kheiri, A., Leibanar, G., Aliramaji, F., Pedram, M., Rezaee, S. and Alighi, M.R. (2015) Rotylenchus castilloi n. sp. (Nematoda: Haplolaimidae) a new spacies with long stylet from northern Iran. Zootaxa 3931(1): 88-100 (doi: org/10.1646/zootaxa.3931.7.6) [4] Rathore, Y.S. and Ali, S.S. (2014) Relationships of root-knot nematode, Meloidogyne incognita, with taxonomic groupings of host plants. Trends in Bioscience 7 (13): 1562-1568. [5] Rathore, Y.S. and Tiwari, S.N. (2016) Relationships of different species of root-knot nematodes to their host taxonomic groupings. Intern. J. Plant Anim. Environ. Sci. 7(1): 29- 36. 1793 Y.S. Rathore

[6] Rathore, Y.S. and Tiwari, S.N. (2016a) Host affinity of nematode species of the genus Aphelenchoides. Inter. J. Sci Res. 5(9): 227-230 [7] Rathore, Y.S. (2017) Host preference in Pratylenchus species. Trends in Bioscience Journal. 10 (32): 6747-6758. [8] Nemaplex (2017) Nemabase host range of genus and species of plant feeding nematodes (retrieved November, 2017). [9] APG IV (2016) Angiosperm Phylogeny Group-An update of the Angiosperm Phylogeny Group classification of orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society, 181 (1):1-20 (doi: 10.1111/boj.12385, retrieved 2016-06-11)). [10] Berneys, E.A. and Chapman, R.F. (1974) Host selection by phytophagous insects (Contemporary topics in entomology), New York, Springer. [11] Hutchinson, J. (1973) Families of flowering plants. 3rd eds., Oxford at the Clarendon Press, 968 p. [12] Subbotin, S.A., Sturhan, D., Vovlas, N., Castillo, P., Tanyi Tambe, J., Moens, M., and Baldwin, J.G. (2007) Application of the secondary structure model of rDNA for phylogeny: D2-D3 expansion segments of the LSU gene of plant parasitic nematodes from the family Haplolaimidae Filipjev 1934. and Evolution 43: 881-890. [13] Tzortzakakis, E.A., Archidona-Yuste, A., Liebanas, G., Birmpilis, I.G., Contalapiedra- Navarrete, C. Navas Cortes, J.A., Castillo, P. and Polomares-Rius, J.E. (2016) Rotylenchus cretensis n. sp. and R.cypriensis Antoniou 1980 at C rete (Greece) with a molecular phylogeny of the genus. European J. Plant Pathol. 144(1):167-184. (Doi: 10.1007/s10658- 015-0760-2).